A riparian plant community from the upper Maastrichtian of the Pyrenees (Catalonia, NE Spain)

Cretaceous Research 56 (2015) 510e529

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A riparian plant community from the upper Maastrichtian of the Pyrenees (Catalonia, NE Spain)
ndez-Marro
n c, Josep Marmi a, *, Carles Martín-Closas b, Mª Teresa Ferna Víctor Fondevilla d, Bernard Gomez e  de Paleontologia Miquel Crusafont, C/ Escola Industrial 23, 08201 Sabadell, Catalonia, Spain Institut Catala Departament d’Estratigrafia, Paleontologia i Geoci encies Marines, Universitat de Barcelona-UB, c/Martí i Franqu es s/n, 08028 Barcelona, Catalonia, Spain c Instituto de Geociencias (CSIC, UCM), Facultad de Geología, C/ Jose Antonio Novais 12, 28040 Madrid, Spain d noma de Barcelona, Avinguda de l'Eix central s/n, 08193 Bellaterra, Catalonia, Spain Departament de Geologia, Universitat Auto e CNRS-UMR5276 LGL-TPE, Universit
e Lyon 1 (Claude Bernard), OSU Lyon, 69622 Villeurbanne, France a

b

a r t i c l e i n f o

a b s t r a c t

Article history: Received 20 March 2015 Received in revised form 18 May 2015 Accepted in revised form 8 June 2015 Available online 7 July 2015

Angiosperms began to colonise riparian habitats very soon in their evolutionary history, probably already in the Aptian, but it is still poorly known when flowering plants finally dominated entirely these kind of
-1) from the upper communities as they do in the present. A new fossil plant locality (Molí del Baro Maastrichtian of the Southern Pyrenees is described in which meandering river facies represent one of the first riparian communities formed only by angiosperms. The fossil assemblage consists of abundant leaves, seeds, logs and sporomorphs. Angiosperms remains dominate in all these cases and the leaf sample is mostly composed of a new eudicot willow-like species, Saliciphyllum gaetei sp. nov., the palm Sabalites longirhachis and an helophytic monocot. Pollen remains suggest that the later belonged to Typhaceae. Most of these plant remains were parautochthonous and deposited in a pond formed in the
-1 represents an unique plant fossil accretional part of a meander loop. The locality of Molí del Baro assemblage in the uppermost Cretaceous of southern Europe. It clearly differs from those reported in other Maastrichtian localities of the Pyrenees (Fumanya and South Isona) and from the CampanianMaastrichtian of Austria and Romania. In addition, it reflects a surprisingly modern physiognomy for a Late Cretaceous riverine plant assemblage that was built up with willow-like plants, palms and reeds. © 2015 Elsevier Ltd. All rights reserved.

Keywords: Riparian community Maastrichtian Angiosperms Pollen Spores Tremp Syncline

1. Introduction The rise of angiosperms to dominate many plant communities and an explosion in their diversity during the Cretaceous changed drastically the plant kingdom, and so affected the whole terrestrial ecosystems. Angiosperms evolved gradually in both time and space, being strongly constrained from the Barremian to the Albian by their ecophysiology and their living environments (Feild et al., 2011; Coiffard et al., 2012). In the Early Cretaceous, European riverine plant communities were rich in conifers, mainly belonging to the Cheirolepidiaceae and the Cupressaceae-Taxodiaceae (Gomez et al., 2001, 2002; Coiffard et al., 2012). As early as the Aptian, these communities began to accommodate early angiosperms in river bank and floodplain environments, but flowering

* Corresponding author. E-mail address: [email protected] (J. Marmi). http://dx.doi.org/10.1016/j.cretres.2015.06.004 0195-6671/© 2015 Elsevier Ltd. All rights reserved.

plants represented only a small part of the whole community in terms of taxonomic diversity. From the Cenomanian to the Campanian angiosperms already formed an important part of riparian communities both in levee and floodplain meandering river environments, alongside with other seed-plants and ferns that still represented one third to half of the assemblage (Coiffard et al., 2012). There is a general agreement that angiosperms dominated all types of environments from lower to middle latitudes during the Maastrichtian. The plant fossil record from outside Europe appeared to be surprisingly rare in this age (Nichols and Johnson, 2008). Nevertheless, this has been beginning to change during the last few years with new data from North America and New Zealand (Spicer and Collinson, 2014). In Europe, continental beds containing Maastrichtian plant micro-, meso- and megafossils have been reported from France, Germany, Poland, Portugal, Romania, and Spain (Petrescu and Dus¸a, 1980; Nichols and Johnson, 2008; Friis et al., 2010; Halamski, 2013). Plant fossils are abundant and

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diverse in the Maastrichtian transitional to continental beds of the Tremp Formation from the northeastern Iberian Peninsula (Mey et al., 1968). The first palaeobotanical studies in localities belonging to the Tremp Formation were conducted during the 1970s and were
dus, 1970, 1972). Palynological mainly focused on sporomorphs (Me works continued during the 1980s (Porta et al., 1985; Ashraf and
dus et al., 1988) and the first attempts to study Erben, 1986; Me the biostratigraphical significance of charophytes were carried out (Babinot et al., 1983; Feist and Colombo, 1983). Since the 1990s to the present, palynological studies of various localities and palaeoenvironments of the Tremp Formation have been focused both on
dus et al., 1992; Mayr et al., 1999; Lo
pez-Martínez miospores (Me
n et al., 2004; Torices et al., 2012; et al., 1999; Fernandez-Marro Villalba-Breva et al., 2012, 2015) and megaspores (Batten et al., 2011). In 1979, the first plant megafossil assemblages were
n discovered near the village of Isona, in the Tremp Syncline (Barro
guez, 1992; Vicente i Castells, 2002). The plant community and Die that lived in the coastal wetlands of the Tremp Formation was reconstructed based on charophytes, sporomorphs and plant megafossils collected from the locality of Fumanya, Vallcebre Syncline (Villalba-Breva and Martín-Closas, 2011; Villalba-Breva et al., 2012). Overall, cheirolepidiacean conifers and palms usually dominated, whereas ferns did only locally. By contrast, palynology, palaeobotany and plant taphonomy from the locality of South Isona (Tremp Syncline) suggested that woody angiosperms were diverse and dominated in the fluvial settings of the Tremp Formation, with ferns being abundant in the understory of riparian communities (Marmi et al., 2014; Villalba-Breva et al., 2015), while floodplain ponds were dominated by characeans, especially by species belonging to genus Microchara (Vicente et al., 2015). In the present study we report plant micro-, meso- and mega
fossils collected from the upper Maastrichtian of the Molí del Baro 1 locality, in the southeastern Pyrenees (Catalonia, northeastern Spain). The aims of the study are: i) to reconstruct the plant community based on sedimentology and taphonomy, ii) compare with other coeval localities in terms of composition and diversity, and iii) discuss the evolution of plant communities in fluvial settings from the Upper Cretaceous of Europe. 2. Geological setting The studied area is located to the north of the village of Sant  d'Abella, northeastern edge of the Tremp Syncline, Lleida Roma Province, Catalonia, northeastern Spain (Fig. 1A and B). Shallow
n (Areny) marine, transitional to continental materials of the Are and Tremp Formations are widely exposed in the Tremp Syncline (Mey et al., 1968). They record a marine regression that began at the Campanian-Maastrichtian boundary (Rosell et al., 2001; Riera et al., 2009). The Tremp Formation was divided into four informal units by Rosell et al. (2001), described from the base to the top as follows: (1) a marine to continental transitional Grey Unit mainly composed of grey marls, lignites, charophyte limestones and sandstones; (2) a fluvial Lower Red Unit with red lutites, sandstones and palaeosols; (3) the lacustrine Vallcebre Limestone and laterally equivalent strata; and (4) the fluvial Upper Red Unit consisting of red lutites, sandstones, and conglomerates. In the Tremp Syncline, palaeontological and biostratigraphic data suggest a Maastrichtian age for the Grey and Lower Red Units of the Tremp Formation (Riera et al., 2009 and references therein; Villalba-Breva and MartínClosas, 2013).
-1 corresponds to the upper part of The locality of Molí del Baro the Lower Red Unit, which was deposited in a fluvial environment with limited tidal influence (Rosell et al., 2001; Riera et al., 2009; Díez-Canseco et al., 2014). The general stratigraphic succession

511

consists of sandstones representing meandering channels interbedded within thick reddish mudstone units (floodplain deposits). The bed containing the plant fossils occurs 70 m below the limestones of Tossal de la Doba and, approximately, 100 m below the
and Suterranya (Riera et al., limestones of Sant Salvador de Tolo 2009), which were attributed to the Palaeocene based on charophyte biostratigraphy by Feist and Colombo (1983). Non-reworked planktonic foraminifera confirmed the Palaeogene age of these limestones (Díez-Canseco et al., 2014). Vila et al. (2012) placed the
-1 in the 29r magnetochron, resulting in a late Molí del Baro Maastrichtian age (Fig. 1C). The studied section belongs to a major, 10e15 m-thick, sandstone body divided into two units separated by a reactivation surface (Fig. 2A). These lower and upper units evidence two main episodes of river activity. The whole sandstone body exhibits a fining-upward trend ranging gradually from coarse sandstones to siltstones and mudstones. Plant bioturbation is extensively developed as colour mottling and often obliterates sedimentary structures. When visible, often in the lower parts of an accretion bar, they appear as cross-laminations followed by current ripples to the top of each bar (Fig. 2A and B). The channel architecture is determined by normally gradeddfrom sandstones to mudstonesdlateral accretions configuring inclined heterolithic stratification (IHS, Fig. 2A), a feature observed in meandering rivers (Thomas et al., 1987; Miall, 1996) and also described in other sandstone bodies of the Tremp Formation (Díez-Canseco et al., 2014). Interbedded between the accretion surfaces, marly deposits (about 45% of CaCO3) yield the main fossiliferous beds (i.e., beds 2 and 3 in Fig. 2C and D). Bed 2 consists of a 20 m-long marl that is 0.3e1 m thick at the top of the lower sandstone unit (Fig. 2A and B). Sandy lenses (bed 3), around 8e15 cm thick but which exhibit limited lateral continuity, appear within these fine sediments. Their contacts are diffuse, passing gradually to the adjacent marls, and no erosive bases are observed. The deposits of bed 3 contain more quartz grains than the marls of bed 2. The major coarse components of the lenses are intraclasts such as edaphic nodules (both of carbonate and oxide). They also contain oncoids and coated grains, as well as fewer rhizolith fragments, charophytes and planktonic foraminifera. In addition to plants, the marls (bed 2) and the coarser level (bed 3) yielded vertebrate, arthropod and mollusc remains (work in progress). Based on all this evidence, the sediments of Molí
-1 are interpreted as the inner, non-erosive part of a del Baro meander loop recording periods of low energy flow, with accumulation of mud in the meander margins, alternating with periods of higher energy flow, with accumulation of sands on the lateral accretion surfaces. The lenses within the marls may be the result of local sedimentary processes, not necessarily linked to the current activity (e.g. collapse of the river margins, floodplain sediment arrivals due to rainfall). 3. Materials and methods
-1 was conducted Fieldwork at the locality of Molí del Baro during the summers of 2002, 2007, 2010 and 2011. One hundred and eighty-five hand rock specimens with leaves, stems and seeds were collected from beds 2 and 3 (Fig. 2B) and are currently housed in the collection of the Museu de la Conca Dell a (MCD) (Isona i Conca Dell a, Lleida, Catalonia, Spain). In addition, pictures were taken of a large log and of sixteen palm leaves, which could not be collected during the excavation. Detailed stratigraphy of the locality was carried out in 2010 and 2011 and the exact stratigraphic location of specimens (n ¼ 108) was determined during these two campaigns. Plant fossils were photographed with a Panasonic DMC-FZ18 digital camera and details were examined and photographed

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-1. A, Geological map of the south-central Pyrenees, showing the localition of the Cretaceous-Palaeocene Tremp Formation, Fig. 1. Geological context of the locality of Molí del Baro
pez-Martínez and Vicens (2012). B, Geological map of the Isona area (enlarged from A) with the position of the Molí del Baro
section (MB) and other correlated modified from Lo
-1. Charophyte biostratigraphic data and magnetostratigraphy are taken after Feist sections (I, and II), modified from Riera et al. (2009). C, Stratigraphic framework of Molí del Baro and Colombo (1983) and Vila et al. (2012), respectively. Section MB, I and II correspond to sections XXIII, XXIV and XXV of Riera et al. (2009).

under a Leica M60 stereomicroscope with integrated digital camera. Drawings of leaves and details of veins, as well as measurements, were made from digital pictures, the latter using the freeware ImageJ v. 1.40 (Rasband, 1997e2008). Angiosperm leaves were described and morphotyped following the methods of Ellis et al. (2009). Plant megafossils were compared with Cretaceous plants known in Europe and North America including the Upper Cretaceous of Nebraska and Kansas, USA (Newberry, 1898; Upchurch and Dilcher, 1990), the Albian-Maastrichtian of Portugal (de Saporta, 1894), the Cenomanian-Senonian of Bohemia, Czech Republic (Kva cek, 1983, 1992; Uli cný et al., 1997; Kvacek and
chov
V
achov
a, 2006; Va a and Kvacek, 2009), the upper Santonian of Fuveau, France (de Saporta, 1890), the lower Campanian of Grünbach, Austria (Herman and Kva cek, 2010), the CampanianMaastrichtian of the Pomeranian-Warsaw-Lubin-L'viv and Use
d

w synclines in Poland and western dom-Szczecin-Lo z-Miecho Ukraine (Halamski, 2013), the lower Maastrichtian of Isona, in Catalonia, Spain (Vicente i Castells, 2002; Marmi et al., 2014) and  and Hat¸eg basins in Romania the Maastrichtian of Rusca Montana (Popa et al., 2014). In addition, pictures of cleared extant leaves from the collections of the Smithsonian Institution (National Cleared Leaf Collection-Wolfe (NCLC-W)) (Das et al., 2014) were used for comparison. Seeds were compared with specimens from the Upper Cretaceous of Europe described by Knobloch and Mai (1986), as well as with extant seeds with botanical affinity to
-1 (e.g. palms). some taxa described in Molí del Baro One sample of grey marls (around 4 dm3) from bed 2 was processed for sporomorph analyses. The sample was prepared

following established procedures of crushing, treatment by hydrochloric (HCl) and hydrofluoric (HF) acids, oxidation to clear organic debris associated and to lighten the colour of palynomorphs. Sieving was performed with a sieve of 10 mm mesh size. Slides were mounted in glycerine jelly and examined under a Leitz Laborlux D light microscope. Photomicrographs were taken with a Wild MPS 41/51 S at the Instituto de Geociencias (CSIC-UCM) of Madrid. The palynological assemblage was analysed and compared with palynofloras of similar age (Kedves and Herngreen, 1980; Herngreen et al., 1986) and reported in the Maastrichtian from
dus, 1970, 1972; Porta et al., 1985; Ashraf the southern Pyrenees (Me
dus et al., 1992; Mayr et al., 1999; Lo
pezand Erben, 1986; Me
n et al., 2004; Torices Martínez et al., 1999; Fernandez-Marro et al., 2012; Villalba-Breva et al., 2012, 2015).

4. Systematic palaeontology 4.1. Plant megafossils 4.1.1. Ferns Clade Monilophyta. Order Unknown. Family Unknown. Genus Sphenopteris (Brongniart) Sternberg. Sphenopteris sp. Fig. 3A. Material: MCD5467.

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-1. A, Picture of the Molí del Baro
locality. B, Scheme of the fluvial architecture and sedimentology of the Molí del Baro
showing the Fig. 2. Sedimentological context of Molí del Baro major structures. C and D, Detailed sections (marked in B) showing the fossil beds 2 and 3 excavated in the summers 2002, 2007, 2010 and 2011.


-1. A, Apex of a frond assigned to Sphenopteris, MCD5467. B, Once pinnataely compound leaf Fig. 3. Fern and conifer remains from the upper Maastrichtian of Molí del Baro attributed to an indeterminate fern, MCD 5249. C, Isolated needle-shaped conifer leaf showing a prominent medial groove (arrowed), MCD5502.

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Description: The single specimen consists of a distal part of a frond. It is 2.1 cm long and 9.4 cm wide. Distal pinnae are completely fused forming a lamina with dentate margins. This lamina is symmetrical and the apex acute. Below the apical lamina, pinnae are entire margined and fused at their bases. The primary vein is straight and tapers towards the apex. Secondary veins arise from the primary vein at acute angles (13 23 ) and are once or twice dichotomously ramified. Comparison: The studied specimen resembles Sphenopteris ungerii Kva cek and Herman from the lower Campanian of Grünbach (Austria) based on the fusion of pinnae at the distal end of the frond and sphenopteroid venation (Herman and Kva cek, 2010: pl. 7, 1e2). However, this species was described from bipinnate fronds, a feature that is not visible in the studied specimen. The genus Sphenopteris is used in a broad sense. It includes sterile foliage of fern-like fronds with basally fused pinnules and showing pinnate venation with lateral veins arising from the midvein at acute angles and dichotomizing a few times (Arnold, 1947; Herman and Kva cek, 2010). Monilophyta gen. et sp. indet. Fig. 3B. Material: MCD5249, 5505, 5537. Description: The most complete specimen (MCD 5249, Fig. 3B) consists of a once-pinnataely compound leaf fragment with opposite leaflets. It is 32.6 mm long and 24.8 mm wide. The best preserved leaflet is 11.8 mm long and 5.3 mm wide. It is symmetrical and has toothed margins. Teeth are regularly spaced and straight/ convex or convex/convex. The sinuses are angular. There is one order of teeth. A single primary vein is present in some leaflets. MCD5537 is an isolated leaflet that is 18.2 mm long and 7.3 mm wide. Comparison: The lack of venation details prevents elucidating if these specimens correspond to fern or angiosperm foliage. However, Cretaceous pinnate foliage is usually assigned to ferns. On this basis, the specimen is tentatively classified as an indeterminate fern. 4.1.2. Conifers Order Unknown. Family Unknown. Coniferophyta gen. et sp. indet. Fig. 3C. Material: MCD5502e3. Description: The specimens are isolated needle-shaped leaves. The MCD5502 (Fig. 3C) is 3.5 cm long and 0.1 cm wide. It has a 0.4 mm wide, prominent groove lengthwise to the medial axis. It is probably the central vascular bundle. Comparison: Conifer twigs with long, needle-shaped leaves from the Cretaceous of Europe are usually assigned to the genus Cunninghamites Presl in Sternberg or Geinitzia Endlicher (e.g. Herman and Kva cek, 2010; Bosma et al., 2012; Halamski, 2013). However, in these genera, leaves are always found attached to twigs or branches. The studied specimens consist of isolated leaves and lack important characters (e.g. leaf arrangement, presence and shape of leaf cushions), which are necessary for a precise identification. 4.1.3. Angiosperms Clade Monocots. Order Unknown. Family Unknown. Monocot type 1. Fig. 4A and B. Material: MCD5251, 5260, 5327, 5339, 5342e4, 5425, 5453, 5510.

Description: The specimens consist of middle fragments of ribbon-shaped leaves with parallelodromous venation. They are 10.0e78.1 mm long and 3.1e18.6 mm wide. The margins are entire and unarmed (i.e. lacking spines). Two longitudinal vein orders are distinguished. Narrow veins (around 0.05 mm wide) are intercalated between thicker primary veins (0.14e0.20 mm wide) (Fig. 4B). Thinner transversal veins can also be observed (Fig. 4B). Comparison: Leaf fragments with the same venation pattern were described in the locality of South Isona by Marmi et al. (2014:
-1 are wider. Ribpl. II, 7). However, specimens from Molí del Baro bon shaped leaves with parallelodromous venation are common from the Cretaceous of Austria, France and Portugal, but intercalated thinner veins are absent (e.g. de Saporta, 1890: pl. II, 2, 4; de Saporta, 1894: pl. 10, 22) or their number is higher than one (e.g. de Saporta, 1890: pl. II, 5; de Saporta, 1894: pl. XXII, 8; Kva cek and Herman, 2004: pl. IX, 1e2). Thus, they are unlike the studied specimens. Monocot type 2. Fig. 4C. Material: MCD5471. Description: A small leaf fragment with single order of longitudinal veins. It is 16.1 mm long and 6.3 mm wide. Longitudinal veins are 0.1 mm wide. Distances between these veins are 0.33e0.54 mm, yielding a density of 22e30 veins per cm. Narrower veins (~0.05 mm wide) are transversal or slightly oblique to primary veins (Fig. 4C). Comparison: Rhizocaulon subtilinervium de Saporta and Typhacites rugosus de Saporta from the upper Santonian of Fuveau, France (de Saporta, 1890: pl. II, 2, 4) have the same vein pattern of MCD5471. These features were also observed in a few leaf fragments from the lower Maastrichtian of Fumanya, in the neighbouring Vallcebre Syncline (Fig. 5f and g in Villalba-Breva et al., 2012). These specimens and specimens from the localities of Isona were tentatively assigned to Pandanaceans (Villalba-Breva et al., 2012; Marmi et al., 2014). However, the studied specimen is assigned to unidentified monocot foliage due to its fragmentary nature and the lack of diagnostic features such as the presence of spines on the margins and an M-shaped transverse section. Order Pandanales. Family Pandanaceae Brown. Genus Pandanites Tuzson. Pandanites sp. Fig. 4D and E. Material: MCD5526. Description: A single specimen consists of a fragment of ribbonshaped leaf (Fig. 4D). It is 24.1 cm long and 3.1 cm wide. Up to 11 spines arise from the same side of the lamina at acute angles (35.5 e42.9 ), pointing to the apex of the leaf. The largest spine is 10.1 mm long and the base 4.7 mm wide (Fig. 4E). There are 1e2 spines per cm. The venation pattern consists of single order, parallel longitudinal veins (Fig. 4E). Distances between veins are 0.3e0.8 mm, yielding a density of 12e30 veins per cm. Narrower veins are transversal to slightly oblique. Comparison: The following diagnostic characters of the genus Pandanites were observed in the studied specimen: elongated, linear leaves, parallel veins of one order, transverse veins thin and marginal spines (according to Kva cek and Herman, 2004). However, other features such as M-shaped transverse section, prominent midrib or auriculate base are not preserved in MCD5526. The vein spacing is consistent with the range given by Kva cek and Herman (2004) for P. trinervis (Ettingshausen) Kva cek and Herman from the lower Campanian of Grünbach (Austria). However, the studied specimen has large marginal spines similar to those reported in P. spinatissimus Petrescu et Dus¸a emend. Popa et al.

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-1. A, Monocot type 1, fragment of leaf showing parallelodromous venation, MCD5453. B, Monocot type 1, detail of Fig. 4. Monocots from the upper Maastrichtian of Molí del Baro specimen MCD5327 showing two orders of parallel longitudinal veins and thin transversal veins. C, Monocot type 2, detail of specimen MCD5471 showing a single order of parallel longitudinal veins and thin transversal veins. D, Fragment of a Pandanites sp. leaf bearing spines at the margin, MCD5526. E, Detail of spines and venation. F, Impression of a lamina of Sabalites longirhachis bearing a prominent costa, MCD5263. G, Impression of an S. longirhachis costapalmate leaf bearing the petiole attached, scale bar equals to 20 cm.

 (Popa et al., 2014: pl. V, from the Maastrichtian of Rusca Montana 2e3). Order Arecales. Family Arecaceae Schultzenstein-Schultz. Genus Sabalites de Saporta. Sabalites longirhachis (Unger) Kva cek et Herman. Fig. 4F and G. Material: MCD5248e9, 5263e65, 5268, 5275, 5277, 5324e5, 5328, 5330, 5332e4, 5416, 5418e20, 5491e2, 5529. Description: Most of remains (n ¼ 20) consist of fragments of leaf laminas with parallel, induplicate and fused segments, sometimes

arising from partially preserved costae. Other specimens have almost complete laminas, composed of fused segments with parallelodromous venation that rise from long costae at acute angles of 25 e47 (Fig. 4F and G). The most complete specimen (Fig. 4G) has a large lamina (51.4 cm long and 31.7 cm wide) attached to a long petiole (28.9 cm long and 4.4 cm wide). The costa is elongated (22.5 cm), massive (4 cm at the base) and narrows gradually towards the apex. Comparison: Large costae bearing laminas with fused segments are typical of the palm leaves of Sabalites longirhachis as reported by Kvacek and Herman (2004) and Marmi et al. (2010). This species is

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-1. A, Complete leaf, MCD5433. B, Middle part of a lamina, MCD5443. C, Apical end of a lamina with well-preserved broFig. 5. Saliciphyllum gaetei sp. nov. from Molí del Baro chidodromous venation, MCD5256. D, Basal end of a lamina with a short petiole attached, MCD5460. E, Fragment of a lamina showing eucamptodromous secondary veins and percurrent tertiary veins, MCD5338. F, Fragment of a lamina showing well-preserved tertiary to quinternary veins, MCD5472. G, Fragment of lamina showing eucamptodromous and looped secondary veins, MCD5262. H, Detail of eucamptodromous secondary veins and percurrent tertiary veins, MCD5244. I, Detail of brochidodromous secondary veins and higher order venation, MDC5256.

common in the Upper Cretaceous (upper SantonianeMaastrichtian) of Austria, France, northeastern Iberian Peninsula, and western Romania (de Saporta, 1890; Vicente i

Castells, 2002; Kva cek and Herman, 2004; Villalba-Breva et al., 2012; Popa et al., 2014). Clade Eudicots. Order Unknown.

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-1, interpretative drawings. A, MCD5443. B, MCD5256. C, MCD5338. D, MCD5472. E, MCD5262. F, MCD5244. Fig. 6. Saliciphyllum gaetei sp. nov. from the Molí del Baro

Family Unknown. Genus Saliciphyllum Fontaine. Type species: Saliciphyllum longifolium Fontaine. Saliciphyllum gaetei sp. nov. Figs. 5 and 6. Specific diagnosis: Linear leaves, nanophyllous to microphyllous, entire-margined and bearing a short petiole. Venation pinnate eucaptodromous to brochidodromous. Brochidodromous secondaries forming irregular loops in the apical end. Tertiary veins opposite or mixed percurrent. Quaternary and quinternary veins reticulate with good development of areolation. Etymology: In honour of Rodrigo Gaete Harzenetter, former director of the Museu de la Conca Dell a and who discovered many palaeontological localities in the Tremp Syncline, including the
-1. Molí del Baro Holotype: MCD5443. Paratypes: MCD5244e6, 5250, 5255e6, 5261e2, 5274, 5277, 5326e7, 5336e8, 5340e1, 5346e9, 5426e7, 5433, 5435, 5438e40, 5448, 5460, 5463, 5465, 5468, 5470, 5472e4, 5481, 5484, 5519e21.


-1, to the north of Sant Roma  d'Abella Type locality: Molí del Baro (Isona i Conca Dell a, Lleida Province, Catalonia, Spain) Type horizon: Grey marls within sandstones at the uppermost part of the Lower Red Unit of the Tremp Formation. Its age is late Maastrichtian as determined by magnetostratigraphy. Description: The leaves are petiolate, simple and nanophyllous to microphyllous. Most of laminae are incomplete, linear and symmetrical (Figs. 5 and 6). They are 18.1e75.4 mm long and 3.8e12.1 mm wide. The petioles are short and marginally attached to the laminae (Fig. 5A and D). They are 2.4e5.1 mm long and 0.7e1.1 mm wide. The margins are entire. The apex is acute and straight (Figs. 5C and 6B), while the base may be acute and straight or slightly decurrent (Fig. 5A and D). The venation is pinnate eucamptodromous, becoming brochidodromous towards the apex (Figs. 5B and C, EeI; 6). The midvein is stout and straight. Secondary veins are thin, irregularly spaced and arise from the midvein at acute angles (29.2 e49.4 ). Brochidodromous secondary veins form irregular loops (Figs. 5C, I; 6B). The intercostal and epimedial tertiary veins are opposite to mixed percurrent and form obtuse

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angles with the midvein (Figs. 5EeH; 6). Quaternary and quinternary veins are reticulate, with well-developed areoles (Figs. 5F, I; 6BeF). Comparison: Lanceolate to linear, entire-margined leaves with pinnate brochidodromous venation are frequent from the Upper Cretaceous of Europe (Kvacek, 1992; Herman and Kvacek, 2010; Marmi et al., 2014 and references therein). Many such leaves
ritier were assigned to genera Myrtophyllum Heer, Eucalyptus L'He de Brutelle, Grevillea Brown, Grevilleophyllum Velenovský and Proteophyllum Velenovský (Kvacek, 1992). These genera are characterised by having intramarginal veins that are lacking in the studied specimens. In the neighbouring locality of South Isona (lower Maastrichtian, Tremp Syncline), microphyllous, linear and entiremargined leaves with pinnate eucamptodromous venation are abundant (Vicente i Castells, 2002; Marmi et al., 2014; VillalbaBreva et al., 2015). They were assigned to a single morphotype (dicot type 3) by Marmi et al. (2014). Some specimens from South Isona (e.g. MGB38383, 38386) have intercostal, percurrent tertiary veins, while in others the intercostal tertiary vein fabric appears

composite admedial (e.g. MGB38388, Marmi et al., 2014: pl. III, 3;
-1 are very similar in size IV, 3). The linear leaves from Molí del Baro and shape to Dicotylophyllum sp. 7 from the lower Campanian of Grünbach (Herman and Kva cek, 2010: pl. 25, 8e14; fig. 53 in Herman and Kva cek, 2010). Both leaf forms have pinnate camptodromous venation. However, in Dicotylophyllum sp. 7, secondary veins arise at very acute angles, running almost parallel to the midvein (Herman and Kva cek, 2010: pl. 25, 11). Some species within the extant genus Grevillea Brown (Proteaceae) have small,
-1 (e.g. G. linearis Brown, linear leaves like those from Molí del Baro UC1089545 in the NCLC-W database, or G. oleoides Sieber, UC249342 in the NCLC-W database). These species have also a stout and straight primary vein but revolute margins and thin secondary veins terminating at the intramarginal vein. Several species within the genus Salix Linnaeus have linear to lanceolate leaves with entire margins and secondary to quinternary venation closely resembling the studied specimens: Salix argophylla Nuttal (UC1221893 in the NCLC-W database) has brochidodromous secondary veins irregularly spaced like MCD5256 (Figs. 5C and 6B) and S. viminalis


-1. A, cf. Cocculophyllum, MCD5506. B, Dicotylophyllum sp. A, MCD5479. C, Dicotylophyllum sp. B, MCD5417. D, Dicotylophyllum sp. C, Fig. 7. Eudicot-like leaves from the Molí del Baro MCD5254. E, Dicotylophyllum sp. D, MCD5253. F, Dicotylophyllum sp. E, MCD5249.

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-1, interpretative drawings. A, cf. Cocculophyllum, MCD5506. B, Dicotylophyllum sp. A, MCD5479. C, Dicotylophyllum sp. B, MCD5417. Fig. 8. Eudicot-like leaves from the Molí del Baro D, Dicotylophyllum sp. C, MCD5254. E, Dicotylophyllum sp. D, MCD5253. F, Dicotylophyllum sp. E, MCD5249.

Linnaeus (UC599331 in the NCLC-W database) has eucamptodromous secondary veins, tertiary veins percurrent and quaternary and quinternary veins reticulate like MCD5244 (Figs. 5H and 6F). The leaves of the olive willow (Salix elaeagnos Scopoli) are also very similar in size, shape and venation to fossil leaves from Molí del
-1. In this sense, they are tentatively attributed to eudicots and Baro assigned to the genus Saliciphyllum which was erected by Fontaine (1889). Order Unknown. Family Unknown. Genus Cocculophyllum Velenovský. cf. Cocculophyllum. Figs. 7A and 8A. Material: MCD5506. Description: The single specimen consists of a fragment of a probably entire-margined lamina. It is 27.1 mm long and 14.8 mm wide. The venation seems pinnate. The midvein is stout. Secondary veins are long, widely spaced and run parallel to the margin of the lamina. Tertiary veins are percurrent and perpendicular to secondary veins. The epimedial tertiary veins are proximally acute to the midvein and distally parallel to intercostal tertiary veins. Quaternary and quinternary veins are reticulate, forming welldeveloped areoles. Comparison: The specimen MCD5506 clearly resembles dicot type 6 from the lower Maastrichtian of South Isona based on size,

entire-margins and features of secondary to quinternary venation (Marmi et al., 2014: pl. III, 6; IV, 6). Although dicot type 6 is imperfect acrodromous, it is not clear whether MCD5506 is pinnate or acrodromous because its basalmost and apical ends are not preserved (Figs. 7A and 8A). However, the possibility that MCD5506 and dicot type 6 belonged to the same taxon cannot be ruled out. Leaves with acrodromous venation and percurrent tertiary veins are found in extant Magnoliids (e.g. Piperaceae [Arctottonia yucatanensis Trelease, MO1084903 in the NCLC-W database] and Laur~ g and Merrill, aceae [Cinnamomum trichophyllum Quisumbín UCH329864 in the NCLC-W database]) as well as Eudicots (e.g. Rhamnaceae [Ampelozizyphus amazonicus Ducke, MO1262968 in the NCLC-W database] and Myrtaceae [Rhodamnia trinervia Blume, UCH267488 in the NCLC-W database]). However, in most of these species, lateral primary veins go beyond the middle of the lamina and form convergent arches towards the leaf apex unlike dicot type 6. Menispermaceae includes some species that are imperfect acrodromous with percurrent tertiary and reticulate quaternary and quinternary veins that closely resemble MCD5506 and dicot type 6 (e.g. Hyperbaena hendunonsis Standley, UCH519161 in the NCLC-W database, and Albertisia laurifolia Yamamoto, uncoded in the NCLC-W database). Leaf fossils assigned to genera Menispermites Lesquereux, Menispermophyllum Velenovský and Cocculophyllum Velenovský have been compared to extant Menispermaceae (Friis et al., 2011). The venation of the studied

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specimen resembles C. cinnamomeum Velenovský from the Cenomanian of Bohemia (cf. fig. 1N in Kva cek, 1983). In this sense, MCD5506 is tentatively assigned to cf. Cocculophyllum even though, based on cuticular features, Kva cek (1983) pointed out that this genus resemblances more to Lauraceae than to Menispermaceae. Order unknown. Family unknown. Genus Dicotylophyllum de Saporta. Dicotylophyllum sp. A. Figs. 7B and 8B. Material: MCD5479. Description: The specimen is a microphyllous, partially preserved leaf, 40.7 mm wide. The apical and basal ends are lacking. The leaf blade is symmetrical and probably entire-margined. The primary vein is pinnate and stout. The secondary veins are probably arcuate and arise from the midrib at wide angles. They are thinner and connect with each other forming loops. Intercostal tertiary veins are sinuous, chevroned or alternate percurrent. Exterior tertiary veins are variable, some of them terminating at the margin while others are looped. Comparison: Based on venation, MCD5479 resembles Populus cyclophylla Heer from the Cretaceous of Dakota Group, Nebraska and Kansas, USA (cf. Newberry, 1898: pl. IV, 1). This species has also entire-margined blades. Some living species within the genus Populus Linnaeus (e.g. P. tremuloides Michaux UCH351862 in the NCLC-W database) have similar vein features: secondary veins arcuate and looped, percurrent intercostal tertiary veins that may be sinuous, chevroned or alternate, as well as exterior tertiary veins terminating at the margin or looped. However, this type of venation is not exclusive to this salicacean genus. It has also been observed in fossil leaves from the Upper Cretaceous described as trochodendroid-like forms (e.g. fig. 22 in Tomsich et al., 2010). Nevertheless, trochodendroid leaves usually have crenate margins. Based on its fragmentary nature and the uncertain taxonomic affinities, the specimen is assigned to the genus Dicotylophyllum de Saporta, which is usually used as a formal name for poorly preserved fossil leaves which differ significantly from other known fossil taxa (Herman and Kvacek, 2010). Dicotylophyllum sp. B. Figs. 7C and 8C. Material: MCD5240, 5417. Description: The best preserved specimen (MCD5417) is a fragment of a microphyllous leaf that is 29.1 mm long and 28 mm wide (Figs. 7C and 8C). Leaf margins are not preserved but the leaf base appears cordate. The venation is actinodromous with up to 6 veins arising from a point near the base. The midvein is straight while accessory primary veins and secondary veins are arched. Intercostal tertiary veins are poorly preserved but appear opposite percurrent. MCD5240 is very fragmented but it also shows actinodromous venation with a straight midvein. Comparison: The specimens are similar in size, base shape and primary vein pattern to Menispermites cercidifolius de Saporta from the Albian of Buarcos (Portugal) (de Saporta, 1894: pl. XXXV, 7). However, the latter has alternate percurrent tertiary veins. Extant and fossil species within Cercidiphyllaceae also resemble MCD5417 and have opposite percurrent intercostal tertiary veins (e.g. Cercidiphyllum crenatum Unger (Brown), Kovar-Eder et al., 1998: pl. I, 5; ACL166 in the UCLA-MP database). However, these features are found in species within other eudicot genera such as Populus Linnaeus and Tetracentron Oliver. In this sense, Friis et al. (2011) warned that it may be difficult to reliably assign fossil Cercidiphyllum-like leaf impressions to the extant family Cercidiphyllaceae. Dicotylophyllum sp. C. Figs. 7D and 8D.

Material: MCD5254, 5451. Description: The leaves are simple, nanophyllous, oblong, symmetrical and entire-margined. They are 12.8e14.7 mm long and 4.1e5.0 mm wide, giving a length/width ratio of 2.9e3.2. The apex is acute and acuminate. The midvein is straight, thick and terminates at the apex. Comparison: The specimens resemble Dicotylophyllum sp. 6 from the lower Campanian of Grünbach (Herman and Kvacek, 2010: pl. 33, 4) in gross morphology and midvein thickness. However, Dicotylophyllum sp. 6 is up to 8 times larger than the specimens
-1. from Molí del Baro Dicotylophyllum sp. D. Figs. 7E and 8E. Material: MCD5253. Description: The leaf is simple and probably microphyllous. It is 54.5 mm long, 28.5 mm wide and only preserves the central part. The venation is pinnate. The midvein is straight. Secondary veins are alternate and excurrently attached to the midvein. They arise from the midvein at acute angles (44.3 e62.2 ). Comparison: Leaves with monopodial venation are common in the Maastrichtian from the southern Pyrenees. However, additional characters such as secondary and tertiary vein frameworks as well as blade marginal features are necessary to compare them with given species or leaf morphotypes. Dicotylophyllum sp. E. Figs. 7F and 8F. Material: MCD5247, 5249, 5258. Description: The leaves are simple, nanophyllous to microphyllous, likely elliptic, symmetrical and entire-margined. They are 17.4e31.0 mm long and 7.9e9.3 mm wide. The lamina base is acute and straight. Venation is pinnate. The primary vein is straight and stout and has excurrently arising secondary veins. Comparison: Elliptic to linear leaves of unknown botanical affinity from the lower Campanian of Grünbach were assigned to several species within Dicotylophyllum de Saporta by Herman and Kva cek (2010). A more precise identification for specimens from
-1 is avoided because the lack of secondary and Molí del Baro higher-order vein details.

4.2. Plant mesofossils Carpolithes sp. 1. Fig. 9AeC. Material: MCD5252, 5269, 5432, 5434, 5437a, 5444, 5466, 5469, 5489, 5504, 5508, 5523, 5525. Description: The seeds are mostly oval or elliptic and flattened. They are 7.0e34.1 mm long and 4.3e15.4 mm wide. The length/ width ration is 1.2e2.2. They are covered by a single, coaly envelope, the surface of which is smooth (Fig. 9A). In some specimens, there is a long and sinuous groove parallel to the outline (Fig. 9B and C). Comparison: The specimens are covered by a single integument, which suggests affinity to gymnosperms (Batten and Zavattieri, 1996). Following Eriksson et al. (2000), the volume of the studied seeds (47.5e2916.7 mm3) is clearly higher than that estimated for angiosperm seeds from the Upper Cretaceous. Thus, the botanical affinity for Carpolithes sp. 1 appears closer to conifers than to angiosperms. Conifers are poorly represented in the pollen sample
-1 (see below). However, pollen attributed to from Molí del Baro different types of conifers was previously reported from the neighbouring Maastrichtian localities of Fumanya and South Isona (Villalba-Breva et al., 2012, 2015). Carpolithes sp. 2. Fig. 9DeF.

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-1 site. AeC, Carpolithes sp. 1 MCD5252, 5444, 5504, respectively (arrows mark the groove). DeF, Carpolithes sp. 2, MCD5350a, 5266aeb, Fig. 9. Seeds from the Molí del Baro respectively. GeI, Carpolithes sp. 3, MCD5482 (micropylar pole, lateral view, and proximal end respectively). JeK, Carpolithes sp. 4, MCD5513b (lateral and basal views respectively). LeM, Carpolithes sp. 5, MCD5350b (lateral and basal views, respectively). NeO, Carpolithes sp. 6, MCD5350c (lateral and apical views respectively. PeT, Carpolithes sp. 7, MCD5516, attributed to Arecaceae (lateral views (PeR) and opposite ends (SeT)). Abbreviations: h, hilum; m, micropyle; r, raphe; vb, vascular bundle.

Material: MCD5266aeb, 5267, 5350a, 5421, 5442, 5449e50, 5477e8, 5485, 5493e6, 5498e5500, 5505aeb, 5513ae5, 5524, 5531. Description: The seeds are ellipsoid to spherical, 3.6e12 mm long and 2.1e8.6 mm wide. The length/width ratio is 1e1.8. Most specimens are covered by a brownish envelope (Fig. 9DeE). The surface is smooth or wrinkled. The hilum is rounded (1.0e2.2 mm in diameter) or elliptic (0.7e1.5 mm long, 0.5e0.9 mm wide) (Fig. 9F). A long and sinuous groove connects with the hilum (Fig. 9E and F). Comparison: The longitudinal groove is interpreted as the raphe. Thus, Carpolithes sp. 2 likely developed from an anatropous ovule. In extant angiosperms, anatropous ovules are most probably ancestral, but curved ovules are also found in some gymnosperms (Endress, 2011). Taking into account the abundance of flowering plants in

the megaflora and sporomorph samples, the Carpolithes sp. 2 is tentatively assigned to an angiosperm of uncertain botanical affinity. Carpolithes sp. 3. Fig. 9GeI. Material: MCD5267, 5454, 5482, 5522, 5531. Description: The seeds are hemispherical and covered by a coaly envelope (Fig. 9G and H). They are 3.5e5.6 mm in diameter. The surface is smooth or slightly rough. Two central concavities are located in opposite poles (Fig. 9G and I). The smaller concavity is 1.2e1.7 mm in diameter and is interpreted as the micropylar end (Fig. 9G). The larger concavity, at the opposite end, is interpreted as the hilum (Fig. 9I). It is 1.7e2.0 mm in diameter and, in some cases, preserves remains of vascular bundles (Fig. 9I). Comparison: The micropyle and the hilum are at opposite ends and the lack of raphe suggest that the seeds developed from an

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orthotropous ovule. The circular depression at the micropylar end might indicate that the seed was operculate. It resembles Discosemen nanum Knobloch and Mai from the Maastrichtian of Eisleben (Germany) (Knobloch and Mai, 1986: pl. XXXII, 32e35). However, this species is smaller in size and developed from anatropous ovules. Carpolithes sp. 4. Fig. 9J and K. Material: MCD5513b. Description: The specimen is reniform, 5.0 mm long and 3.7 mm wide. The length/width ratio is 1.3. It is covered by a coaly envelope with smooth surface. In its concave side, it has a bulge 1.2 mm long and 0.6 mm high. It is most likely the vascular bundle. The outline of the hilum is not preserved. Next to the bulge, there is a longitudinal crest that is 2.7 mm long and 0.4 mm wide. It might correspond to the raphe. Comparison: Anatropous reniform seeds are found in several taxa within extant Fabales, such as Medicago polymorpha Linnaeus (personal observation). The concave side containing the vascular bundle is interpreted as the hilar notch that is present in members of Fabales as well as in other eudicots such as Solanales. Thus, Carpolithes sp. 4 probably corresponds to an eudicot. Carpolithes sp. 5. Fig. 9L and M. Material: MCD5350b. Description: The single specimen is auriform and covered by a smooth, coaly envelope. It is 2.2 mm long and 1.6 mm wide. The length/width ratio is 1.4. The hilum is oval (0.4  0.3 mm). Right next to the hilum, there is a circular hole (0.31  0.23 mm) that is interpreted as the micropylar region. Comparison: The seed probably developed from an anatropous ovule due to the very close position of the hilum and micropyle. Klikovispermum waltherii Knobloch and Mai from the Maastrichtian of Germany has similar size and a large elliptic to oval trough in one side (Knobloch and Mai, 1986: pl. VII, 28, 33), like the specimen
-1. The specimen also resembles Melisoma from Molí del Baro praealba Knobloch and Mai from the Maastrichtian of Walbeck, Germany (Knobloch and Mai, 1986: pl. XXIV, 18). Both seeds are similar in size and have smooth surface and a concave side surrounded by a rounded edge. However, neither K. waltherii nor M. praealba have circular to elliptic scars close together, which likely represent the hilum and micropylar region in the specimen
-1. from Molí del Baro Carpolithes sp. 6. Fig. 9N and O. Material: MCD5350c. Description: The specimen is spherical and covered by a coaly envelope of rough surface. It is 4 mm in diameter. A prominent crest 0.2 mm wide divides the seed into two hemispheres. It connects with a small circular hole of 0.3 mm in diameter. Comparison: The circular hole is interpreted as the hilum and the crest may be the raphe, suggesting that the development was anatropous. It resembles Laramisemen rothii (Knobloch) Knobloch and Mai from the Upper Cretaceous of central Europe (Knobloch and Mai, 1986: pl. LII,10e12). This taxon is spherical, anatropous and the hilum and micropyle are very close and located at the top of a sharp apical end. The raphe forms a straight crest connecting with the hilum.
-1 is three times larger However, the specimen from Molí del Baro than L. rothii and, in apical view, its outline is oval instead of circular. Genus Laramisemen was attributed to Saxifragales by Knobloch and Mai (1986). Carpolithes sp. 6 also resembles in shape and ornamentation the seeds of some extant Laurales, such as Sassafras albidum

(Nuttall) Nees (personal observation). Species within Sassafras Nees and Ebermaier develop seeds from anatropous ovules. Carpolithes sp. 7. Fig. 9PeT. Material: MCD5437b, 5461, 5483, 5507, 5516, 5535. Description: The seeds are ovoid to ellipsoid and covered by a coaly envelope ornamented with thin longitudinal grooves (Fig. 9PeR). Five thick grooves converge to the sharp micropylar pole giving a pentarradial symmetry in cross section (Fig. 9S and T). In lateral view, the basal end is nearly rounded. The seeds are 16.4e24.9 mm long and 6.5e14.8 mm wide. The length/width ratio is 1.8e2.5. Comparison: Ovoid to ellipsoid seeds bearing a seed coat ornamented with parallel, longitudinal grooves converging in a sharp pole are found in numerous extant palm species (e.g. the foxtail palm, Wodyetia bifurcata Irvine; the umbrella palm, Hedyscepe canterburyana (Moore and Mueller) Wendland; the vine palm, Attalea butyracea (Mutis ex Linnaeus) Wessels Boer). Thus, it is likely that the parent plant of Carpolithes sp. 7 was a member of Arecaceae. 4.3. Plant microfossils The density of grains was low in the collected sample, giving up to 110 sporomorphs from which 29 taxa were identified including ferns and lycophytes (Fig. 10AeE), conifers (Fig. 10FeH), cycadophytes (Fig. 10I), monocots (Fig. 10JeM), eudicots (Fig. 10NeR) and algae (Fig. 10S). Angiosperms dominated the sporomorph assemblage representing the 83% of the sample. Most of them (48%) are assigned to Juglandaceae: Mul
) Thomson and Pflug (Fig. 10N tiporopollenites maculosus (Potonie and O) is reminiscent of Juglans Linnaeus and Polyatriopollenites
) Pflug resembles Pterocarya Nuttall and stellatus (Potonie Moquin-Tandon. Pantoporate grains bearing circular annulus of
) Thomson and Pflug Periporopollenites stigmosus (Potonie (Fig. 10R) comprise the 18% of the sample and are attributed to the genus Liquidambar Linnaeus. In a lower proportion (6%), grains of Polyporopollenites undulosus (Wolff) Thomson and Pflug (Fig. 10Q) are reminiscent of the genus Ulmus Linnaeus. Within monocots, pollen grains attributed to Arecaceae (Mono
) Potonie
and Monosulcites colpopollenites tranquillus (Potonie brevispinosus (Biswas) Sah and Dutta) are frequent (Fig. 10J and K). Some forms related to Typhaceae (Emmapollis sp. and Sparganiaceaepollenites sp.) (Fig. 10L and M) are less abundant. Spores as well as gymnosperm pollens only represent 8% of the sample. Within conifers, pollen grains attributed to Pinaceae (Fig. 10F) as well as taxodioids (Inaperturopollenites concedipites (Wodehouse) Krutzsch and Inaperturopollenites polyformosus (Thiegart) Thomson and Pflug) (Fig. 10G and H) stand out. Fern spores include the families Schizaeaceae and Polypodiaceae (Fig. 10AeC, E). Zygospores of Zygnemataceae (Ovoidites spriggii (Cookson and Dettmann) Zippi) are very scarce (Fig. 10S). 5. Taphonomy 5.1. Plant megafossils
-1, was mainly found The plant fossil assemblage of Molí del Baro in bed 2 and almost entirely consisted of angiosperms (Fig. 11). These specimens included vegetative organs (stems, leaves) (Fig. 12AeD) as well as seeds, the leaves being dominant (69%) (Fig. 11AeC). Eudicot foliage is mostly represented by Saliciphyllum gaetei sp. nov. (Figs. 11DeE, 12DeE). Leaves of Sabalites longirhachis

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-1. A, Verrucatosporites alienus (Monilophyta, Polypodiaceae). B, Klukisporites scaberis (Monilophyta, Schizaeaceae). C, Retitriletes cf. facetus Fig. 10. Sporomorphs from Molí del Baro (Monilophyta, Schizaeaceae). D, Echinatisporis sp. (Lycophyta). E, Chomotriletes fragilis (Monilophyta, Schizaeaceae). F, Pinuspollenites sp. (Coniferophyta, Pinaceae). G, Inaperturopollenites concedipites (Coniferophyta, Cupressaceae). H, Inaperturopollenites polyformosus (Coniferophyta, Cupressaceae). I, Cycadopites carpentieri (Cycadophyta). JeK, Monosulcites brevispinosus (Monocots, Arecaceae). L, Sparganiaceaepollenites sp. (Monocots, Poales). M, Emmapollis sp. (Monocots, Poales). NeO. Multiporopollenites maculosus (Eudicots, Juglandaceae). P, Oculopollis sp. (Normapolles). Q, Polyporopollenites undulosus (Eudicots, Ulmaceae). R, Periporopollenites stigmosus (Eudicot, Liquidambar). S, Ovoidites sprigii (Zygnematacean algae). Scale bar equals to 10 mm.

(17%) and monocot type 1 (11%) are relatively abundant compared to other forms that represent very small portions of the sample (0.5e1.4%) (Figs. 4FeG, 11E, 12C, F). Seeds are diverse and seven types have been distinguished (Fig. 9). Most of them are attributed

to angiosperms. Carpolithes sp. 1 (16%) and 2 (35%) are the most abundant (Fig. 11C). The angiosperm dominance of the plant assemblage is consistent with the general agreement for low to middle latitudes during the Maastrichtian (Coiffard et al., 2012),

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-1. A, Plant megafossils (hand specimens) collected from each bed during the 2010e2011 excavations. Fig. 11. Diagrams showing the abundance of plant remains from Molí del Baro B, Abundance of vegetative organs (stems and leaves) and seeds calculated from the whole sample. C, Abundance seed types from the whole sample. D, Percentages of leaves belonging to major plant groups based on the whole leaf sample. E, Histogram showing the abundance of angiosperm taxa based on leaves. F, Percentages of sporomorphs belonging to major plant groups (spores include ferns and lycophytes and gymnosperms include cycads and conifers). G, Histogram showing the abundance of angiosperm taxa in the sporomorph sample.

though it was not the case in some plant communities in this region (Villalba-Breva et al., 2012). Linear dicot leaves of Saliciphyllum gaetei sp. nov. may form monospecific accumulations in some horizons without any preferential orientation (Fig. 12D). Most of these leaves are wellpreserved, showing limited fragmentation of the leaf lamina, and a few of them are complete bearing petioles attached (Fig. 5A). Alternatively, in other horizons, the leaves of this species may occur isolated or mixed with other leaves. In this case leaf laminae may be very damaged and torn following the secondary veins, displaying a pattern that corresponds to what Ferguson (1985) experimentally obtained when leaves were submitted to high energy hydrodynamic conditions (Fig. 12E). These taphonomic characters suggest

that the leaves of Saliciphyllum gaetei sp. nov. were locally parautochthonous in the depositional setting, forming leaf litter accumulations in ponds of the accretional side of a meander, while other leaves of the same species reached this area from upstream sources after significant transport as suspended load in the river flow. Monocot type 1 leaves are also seen as parautochthonous. They are also relatively abundant and locally accumulated in a pattern similar to Saliciphyllum (Fig. 12C). This species displays leaf morphology compatible with a helophytic plant, which is an ecomorph expected to occur in the depositional setting of leaf bed 2. Palms are also abundant in the fossil assemblage and are represented by leaves (Fig. 4), possible seeds (Fig. 9PeT) and stems (Fig. 12B). Palms are evergreen and the leaves are marcescent (i.e.

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-1. A, Large log from bed 2 excavated during the 2002 campaign (scale bar ¼ 5 cm). B, Fragment of a Fig. 12. Taphonomic features of the plant fossil assemblage from Molí del Baro palm stem (medial section, top; cross section, below) corresponding to the subdermal or central zone with densely packed parallel vascular bundles, MCD5424. C, Randomly orientated leaves of monocot type 1, MCD5510. D, Accumulation of randomly orientated leaves of Saliciphyllum gaetei sp. nov., MCD5341. E, Partial leaf of Saliciphyllum gaetei sp. nov. with tears following secondary veins, MCD5244. F, Fragment of Sabalites longirhachis with other plant remains preserved in different layers, MCD5275.

decaying while still attached to the parent plant). The presence of numerous leaves of S. longirhachis with well-preserved laminae, bearing petioles attached, in the fossil assemblage from Molí del
-1 (Fig. 4FeG) suggests traumatic shedding (e.g. by strong Baro winds) followed by limited transport (Martín-Closas and Gomez, 2004). A similar taphonomic process was inferred for the same species in the coastal wetlands of the Grey Unit at the neighbouring
, which are early Maaslocalities of Fumanya and Coll de Nargo trichtian in age (Marmi et al., 2008, 2010; Villalba-Breva et al., 2012). The scarcity of remains from other species in leaf bed 2 and their poor preservation, with only small leaf portions present, suggests that most of them grew probably far from the depositional setting and reached it after considerable transport and mechanical

fragmentation in the river flow. The few plant fossils found in coarser sandy sediments of bed 3 (mostly seeds) probably correspond to a highly selected assemblage of these organs, easily transported by flotation, before being deposited at the meander loop. Root marks are lacking in the plant bearing beds studied. However, the fluvial sandstone underlying the plant bearing beds is strongly bioturbated and evidence for palaeosols occurs in the underlying and the overlaying beds, which correspond to floodplain facies (Fig. 2) Most plant megafossils were preserved as adpressions suggesting a rapid burial or the lack of contact with air. They were preserved in a fine lutite, strongly limiting percolation by oxidizing meteoric water. Similar features were observed in the locality of

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Fig. 13. Plant palaeoecological and taphonomic reconstruction of the riparian environments from the upper Maastrichtian of southern Pyrenees basin, based on data from Molí del
-1. Red arrows indicate hypothetic origins and transport of parautochthonous plant remains deposited in beds 2 and 3. (For interpretation of the references to colour in this Baro figure legend, the reader is referred to the web version of this article.)

South Isona, in the same basin, which was interpreted as a distal floodplain environment (Villalba-Breva et al., 2015). In spite of preservation of the organic matter as a fine coaly pellicle, the preservation of leaf cuticles is rare or absent, which hinders in most cases a more precise identification of their taxonomic affinities. 5.2. Plant microfossils The studied samples are not rich in sporomorphs and some of them show a poor preservation, suggesting that they were mostly
-1 is dominated by allochthonous. The assemblage from Molí del Baro angiosperms (Fig. 11FeG). Wind-pollinated taxa, including members of Juglandaceae, Altingiaceae and Ulmaceae, are the most represented (Fig. 11G). Conifer pollen and fern spores are scarce. The former are mainly represented by the Taxodiaceae that includes a number of riverine representatives. Fern spores are not abundant (Fig. 11F) also suggesting that the local floodplain vegetation was underrepresented in the sample. There is a very low proportion (less than 10%) of taxa associated with aquatic habitats (e.g. conjugate zygospores and pollen of Typhaceae). These would be the only strictly autochthonous sporomorphs in the river pond sediments. 6. Discussion
-1 has the highest richness The fossil assemblage of Molí del Baro of plant remains known to date from the Maastrichtian localities of the Iberian Peninsula. It includes sporomorphs, a variety of seeds, logs and leaves. Angiosperms are dominant in all cases (Fig. 11), unlike the other Pyrenean Maastrichtian localities known (Fumanya and South Isona), where cheirolepidiacean conifers and ferns, respectively dominated brackish-freshwater swamp and floodplain assemblages together with angiosperms (Villalba-Breva et al., 2012, 2015; Marmi et al., 2014). Even though being abundant, plant


-1 reveal that angiosperm diversity megafossils from Molí del Baro
-1, 90% is low compared to the South Isona locality. In Molí del Baro of the leaf fossils are assigned to three angiosperm taxa (Monocot form 1, Sabalites longirhachis and Saliciphyllum gaetei sp. nov.) while in South Isona S. longirhachis, a possible Pandanites, one indeterminate monocot and up to 15 dicot leaf morphotypes were represented in more similar proportions (Marmi et al., 2014). However,
-1 would be more represenplant megafossils from Molí del Baro tative of local vegetation than those from South Isona, that were largely allochthonous. Taphonomic evidence indicates that the sporomorph sample
-1 was mostly allochthonous and, as a whole, from Molí del Baro represented regional vegetation. Based on their abundance and relatively good preservation, leaves of monocot form 1, Sabalites longirhachis and, especially Saliciphyllum gaetei sp. nov. underwent only short distance transport and their parent plants likely grew up near the depositional setting (i.e. a pond in a meandering river loop). Thus, it is plausible that local vegetation was mainly composed of these three taxa (Fig. 13). Ribbon shaped leaves with parallelodromous veins of monocot form 1 (Fig. 4A and B) might correspond to a member of Typhaceae as it is suggested by the presence of Emmapollis sp. and/or Sparganiaceaepollenites sp. in the pollen sample. This is also consistent with the depositional setting inferred. Most of seed types were likely transported by streams. However, Carpolithes sp. 7 (Fig. 9PeT) closely resembles those of palms and might be produced by Sabalites trees growing near the meander. Previous works suggested that Sabalites longirhachis inhabited the shores of freshwater lakes in the coastal wetlands of the lower Maastrichtian Iberian Peninsula together with the cheirolepidiacean conifer Frenelopsis (Marmi et al., 2010; Villalba-Breva et al., 2012). In the present study, palms have been found associated with other angiosperms (monocots and eudicots) in a fluvial setting

J. Marmi et al. / Cretaceous Research 56 (2015) 510e529

showing limited tidal influence. During the Maastrichtian, emergent lands of the Iberian Peninsula and part of France formed a large island, the so-called Iberoarmorican Domain (Dercourt et al., 2000). Sabalites longirhachis has also been reported from the lower Campanian of Austria and the Maastrichtian of Romania (Kva cek and Herman, 2004; Popa et al., 2014), which were also islands, separated from the Iberoarmorican Domain by the Tethys seawaters. Evidence from Grünbach (Austria) suggests that these extinct palms shared swampy lowlands with pandanaceans and were frequent in wetland juglandaceous forests that included evergreen taxodioid conifers, within a relatively large deltaic plain (Herman and Kva cek, 2010). In Rusca Montan a and Hat¸eg basins (Romania) S. longirhachis is found in volcanic-dominated, fluvial to paralic depositional systems (Popa et al., 2014). All this evidence suggests a remarkable ecological plasticity for this species, which inhabited a wide range of habitats, in wetland and fluvial settings, sharing open to more or less closed plant communities with different plant species and adapted to different climatic conditions, from colder and more humid during the Campanian-early Maastrichtian of Austria and Iberia to hotter and dryer during the Maastrichtian of Romania (Popa et al., 2014).
-1 provides the first unequivocal evidence for panMolí del Baro danaceans in the Maastrichtian Iberian Peninsula. Pandanites-like leaves have been previously reported from the lower Maastrichtian of Fumanya and South Isona (Villalba-Breva et al., 2012; Marmi et al., 2014). However, the only characters that related these specimens with pandanaceans were those of venation, lacking important diagnostic features as the presence of spines and the typical Mshaped transversal section (Kva cek and Herman, 2004). The genus Pandanites is very abundant in the uppermost Cretaceous of Austria and Romania (Herman and Kva cek, 2010; Popa et al., 2014). In the lower Campanian of Grünbach, Pandanites was interpreted as a part of a coastal swamp, coal-forming vegetation (Herman and Kva cek, 2010) that is very different to the environment inferred for Molí
-1. This fact and the presence of only a single individual in del Baro the sample suggests that Pandanites was not a member of the plant community growing near this fluvial setting. Based on plant megafossils, Cenomanian-Campanian riparian plant communities from Europe were dominated by angiosperms including members of Lauraceae and Platanaceae (Coiffard et al., 2007; Coiffard and Gomez, 2010). In the lower Maastrichtian of South Isona, plant megafossils and sporomorphs suggested that members of Betulaceae shared these environments with a variety of ferns, which grew in the understory (Villalba-Breva et al., 2015). The riparian community reconstructed with data from Molí del
-1 differs from those reported in previous studies by its modBaro ern appearance and includes a dominant eudicot species with willow-like leavesdlikely neither related with Lauraceae nor Platanaceae and Betulaceaed as well as monocots (probable Typhaceae and palms). These differences may be due to the true composition of plant communities or taphonomic biases, since plant remains of the fossil assemblage from South Isona were deposited in a distal floodplain setting and were mostly allochthonous. Only one out of fifteen dicot taxa reported in the lower Maastrichtian locality of South Isona was most probably related to channel margin environments based on taphonomic data (VillalbaBreva et al., 2015). 7. Conclusions
-1 (Tremp basin, Eastern The fossil assemblage of Molí del Baro Pyrenees, Spain) would represent one of the first riparian communities growing along meandering rivers entirely formed by angiosperms resembling modern willow-dominated communities.
-1 consists of an unique plant fossil The locality of Molí del Baro

527

assemblage in the uppermost Cretaceous of southern Europe. It is clearly different from those reported from other Maastrichtian localities of the Pyrenees (Fumanya and South Isona) (Villalba-Breva et al., 2012, 2015; Marmi et al., 2014) and from the CampanianMaastrichtian of Austria and Romania (Herman and Kva cek, 2010; Popa et al., 2014). The assemblage included different plant organs: leaves, logs, seeds and sporomorphs. However, the new reconstructed plant community is characterised by a low species richness and grew in a high sinuosity fluvial setting with minor tidal influence. It was likely dominated by a deciduous eudicot resembling extant willows, reed-like monocots and the widespread Sabalites longirhachis palms. The riparian community inferred for
-1 differs from that of the lower Maastrichtian of Molí del Baro South Isona, which was represented by members of Betulaceae and ferns in the understory (Villalba-Breva et al., 2015). Data from Molí
-1 also suggest that, during the late Maastrichtian, particdel Baro ular angiosperms totally dominated the fluvial environments of the Pyrenean basin, some of them, like Saliciphyllum gaetei sp. nov. showing surprising parallelism in their leaf morphology with extant willows that are also frequently growing in riparian communities. As a whole, the reconstructed plant community from
-1 is remarkably similar to many modern angiospermMolí del Baro dominated fluvial margin communities. This suggests that the Cretaceous-Palaeogene (K-Pg) event hardly disrupted the evolutionary trajectory of angiosperm dominated communities in fluvial settings, at least in the southern European archipelago.

Acknowledgements
-1 site were performed under Excavations in the Molí del Baro the permission of the Departament de Cultura i Mitjans de
(Generalitat de Catalunya). This research was also Comunicacio partially funded by projects CGL2011-30069-C02-01/02, CGL201127869, CGL2012-35199 and CGL2013-42643-P of the Spanish Ministerio de Economia y Competitividad and by project SGR2014251 of the Catalan Autonomous Government. V. F. was supported by a FPI grant (BES-2012-052366) from the Ministerio de Economía y Competitividad. We are very grateful to all collaborators that  participated in the digging works and especially to Drs. Angel Galobart, Rodrigo Gaete, Fabio M. Dalla Vecchia and Violeta Riera who directed the fieldworks carried out in this site. We finally acknowledge the valuable comments of an anonymous reviewer, Dr. R.A. Spicer and the editor that improved the manuscript. References Arnold, C., 1947. An Introduction to Paleobotany. McGraw Hill Book Company, New York. Ashraf, A.R., Erben, H.K., 1986. Palynologische Untersuchugen an der Kreide/Terti€ arGrenze west-Mediterraner regionen. Palaeontographica Abteilung B 200, 111e163. Babinot, J.F., Freytet, P., Amiot, M., Bilotte, M., de Broin, F., Colombo, F., Durand, J.P., Feist, M., Floquet, M., Gayet, M., Lange-Badre, B., Masriera, A., Massieux, M.,
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